Revista del Instituto de Investigaciones de la Amazonía Peruana

VERTICAL SEGREGATION IN SPECIES FROM A BAMBOO FOREST IN SOUTHEAST OF AMAZONIA,

1 Jhon Jairo LÓPEZ-ROJAS1 Universidad Nacional de San Martín, Facultad de Ecología. [email protected]

ABSTRACT Pristimantis species was studied in a remanescent forest dominated by bamboo in the southwest region of the In this work, the vertical stratification in

Amazon. The field work was carried out between December 2012 and May were2013. recorded, The differences Pristimantis in height fenestratus of the perch being were the analyzedmost abundant interspecifically, and occurring and intraspecific in relation to the structure of the vegetation. A total of four species height, as well as between adults and juveniles. For P. fenestratus, there was noin alldifferentiation sampling units. of segregation There was between significant sampling interspecific units. The difference structure in of perch the vegetation did not explain the preference of heights; however, leaf litter depth

all sampling units to infer better explanations. influenced its vertical stratification. More data and occurrences are needed in Pristimantis, bamboo forest, structure of the vegetation. KEYWORDS: Vertical segregation,

DISTRIBUCIÓN VERTICAL EN ESPECIES DE PRISTIMANTIS EN UN BOSQUE DOMINADO POR BAMBÚ EN LA REGIÓN SUROESTE DE LA AMAZONÍA, BRASIL

RESUMEN En este trabajo, se estudió la distribución vertical en especies de Pristimantis en un bosque dominado por bambú en la región suroeste de la Amazonía.

Los muestreos se realizaron entre diciembre del 2012 y mayo del 2013. Las Vol. 27 (1) 2018. 47-54 DOI: https://doi.org/10.24841/fa.v27i1.457 47 Vertical segregation in Pristimantis species from a bamboo forest in southeast of Amazonia, Brazil

en relación a la estructura de la vegetación. Un total de cuatro especies fueron registradas,diferencias de Pristimantis la altura de fenestratuspercha fueron fue analizadas la más abundante inter e intra-especificamente ocurriendo en total percha, tanto en adulto como en juveniles. Para P. fenestratus no hubo diferencia delas segregación unidades de entre muestreo. las unidades Hubo una de muestreo. diferencia La significativa estructura en de la vegetación altura de no explicó la preferencia vertical, sin embargo, la profundidad de la hojarasca unidades para inferir mejores explicaciones. explicó su estratificación. Mas datos y ocurrencias son necesarios en todas las Pristimantis, bosque dominado por bambú, estructura de la vegetación. PALABRAS CLAVES: Segregación vertical,

Vol. 27 (1) 2018. 47-54 DOI: https://doi.org/10.24841/fa.v27i1.457 48 Vertical segregation in Pristimantis species from a bamboo forest in southeast of Amazonia, Brazil

INTRODUCTION The dynamics of bamboo growth is crucial in

Niche theory suggests that coexisting species will abundance in this assemblage, differentiating certaininfluencing habitats vegetation of targeted structure species in (López-Rojasrichness and through the segregation of shared resources reduce the effects of interspecific competition In this study the vertical distribution of species often use different tactics without causing a ofet al.,the 2015). genus Pristimantis occurring in syntopy competitive(Chesson, 2000),exclusion these of resources coexisting occupying species in a fragmented forest dominated by bamboo, needs to address questions regarding vertical different habitats (Caramashi, 1981; Hauer and between adults and juveniles. Additionally, does Stanford, 1986; Kiszka et al., 2011). stratification and interspecific differences Several studies have focused vertical segregation of the various species? stratification in Anura (Arroyo et al., 2008; the vegetation structure influence the vertical Muñoz-Guerrero, 2007; Guayasamin and Funk, ecological2009; de activity, Menezes different Gondim, strategies 2013; are Fonseca- used MATERIAL AND METHODS byPérez, the 2017).anuran Tospecies explain to coexist this behavioral in the same and The work was carried out in a forest fragment habitat: ontogenetically (Eterovick et al., 2010, o o de Menezes Gondim, 2013), foraging sites and known as Fazenda Experimental Catuaba femalescompetition during inter the andbreeding intraspecific season by (Duellman strategic (10 04’S, 67 37’W), in the municipality of SenadorGuadua and Pyles, 1983; Toft, 1985), and by attracting weberbaueriGuiomard, Acre State, Brazil. It comprises an area of ​​1200 ha, dominated by bamboo, vocalization frequencies in males (Miyamoto, and analysis of Pilger vegetation (Smith structure and Nelson, variables 2011). are 1982). However, until now there have been no Distribution of plots in Catuaba, measurements approaches that relate vertical intra-specific structuresegregation the (devegetation. Menezes Gondim, 2013) and largelydetailed induring López-Rojas the rainy et al. (2015).season, Field between work interspecificThe focus segregationon anuran species with variablesof the genus that was conducted for a total of 70 days of sampling, Pristimantis involved particularities in their natural history November 2012 and May 2013, with twice of direct development Jiménez de (Pombal la Espada, and 1870, Haddad, has repetitionsmonthly visits, for eacheach plot. five days in length, totaling 180The hours sampling of observations, of Pristimantis with anwas average carried of out 12 at night using active visual and auditory survey 2007), as sensitive indicators of anthropogenic disturbances (Pearman, 1997), and present vertical stratifications in vegetation (Lynch and searching(Crump andthe trail Scott to 1994).an extent Two of two researchers meters theDuellman, most diverse 1997; Guayasamingroups of and Funk, in 2009; the widesurveyed on botheach plotsides for of approximately the central line, 90 minutes, up to a Blair and Doan, 2009). This genus is one of For each individual observed, I recorded the ofNeotropics, the Brazilian with Amazon 38 species is characterized occurring in by Brazil the perchingheight of threeheight meters above betweenground level 17:00 using h - 23:00 metric h. (Segalla et al., 2017). The southwestern region presence of patches of bamboo (Silveira, 2005). measurements. Snout-vent-length (SVL) was Vol. 27 (1) 2018. 47-54 DOI: https://doi.org/10.24841/fa.v27i1.457 49 Vertical segregation in Pristimantis species from a bamboo forest in southeast of Amazonia, Brazil measured with a digital caliper to differentiate P. fenestratus was present in all plots, so it adults from juveniles, under the criteria of Lehr

Likewise,was possible the tovegetation infer if there structure was an variables intraspecific and and Duellman and Lehr (2009), Padial de la difference in all the sampling units by ANOVA. comparisonRiva (2009). with All specimensspecimens encountereddeposited in were the identified in the field or in the laboratory, by heightthe first in axis each of theplot PCA by (see,linear López-Rojas regression. et al.,All Federal do Acre. 2015) were used to relate them to the mean HerpetologicalAfter analysis Collection of homoscedasticity of the Universidade and normality of the data, an analysis of variance analyzes were performed in RStudio 1.0.153 (R, differences 2015) and Statistic 10 (StatSoft Inc., 2011). in the use of the perching height among the species. RESULTS (ANOVA) was performed to identify the of means was calculated by the Tukey-Kramer Perching height above ground level was recorded The significance of the differences between pairs Pristimantis. adults and juveniles was done through a T-test. method. The intraspecific difference between in 229 individuals of four species of There was interspecific difference in relation to

Fig 1. Vertical distribution of four species of Pristimantis in Catuaba, Acre, Brazil.

Vol. 27 (1) 2018. 47-54 DOI: https://doi.org/10.24841/fa.v27i1.457 50 Vertical segregation in Pristimantis species from a bamboo forest in southeast of Amazonia, Brazil

Table 1. Descriptive data and ontogenetic differences in four species DISCUSSION of Pristimantis observed in Catuaba, Acre State, Brazil. N=Number of individuals; M=Mean; SD=Standard Deviation; t=Student t-test; This study showed a more p=Propability. quantitative subject regarding Species N M SD t p P. altamazonicus differencesLynch and Duellman in relation (1997), to andthe Adults 16 35.8 18.2 -2.76 < 0.01 heightit was possible of the to findsubstrate intraspecific used Juveniles 10 69.5 43.1 for these nocturnally active P. fenestratus . Pristimantis exhibit a greater detectability in the rainy Adults 56 22.3 10.0 -3.29 0.00 Juveniles 91 34.2 25.8 P. reichlei presentseason (Arroyowork was et carried al., 2008; out Waldez et al., 2011), and the Adults 15 32.0 23.7 -2.88 < 0.16 month period, in contrast to the Juveniles 13 65.8 35.9 during the majority of this five- P. skydmainus Adults 12 23.8 7.1 -2.3 <0.01 whichwork oftook Miyamoto two months (1982) and two and yearsGuayasamin respectively. and Funk (2009) Juveniles 16 43.0 32.6 Other authors have observed the vertical distribution p of micro-habitats among , and P. altamazonicus P. fenestratus3,225 P. Pristimantis skydmainusthe hanger occupied height (ANOVA, ranges of F heights=8.02, between <0.01, Fig 1). P. reichlei was more species in other regions: western Ecuador (Guayasamani and Funk, 2009; Lynch 20 Perching and 39 cm,height whereas in P. fenestratus was southwestand Duellman, region 1997) of andthe CostaAmazon Rica in(Miyamoto, a forest common between 40 and 59P. cmaltamazonicus (Fig. 1). and dominated1982). Here by I showbamboo. vertical This segregationtype of vegetation in the P. reichlei (Tukey-Kramer method, p presents effects of the dynamics imposed by psignificantly lower than P. skydmainus bamboo, affecting the structure of vegetation (Tukey-Kramer method, p P. reichlei=0.04 and y P.=0.02), skydmanus but didalso not showed differ differences from in perch height (Tukey-Kramer method,>0.05). p vertical(Griscom segregation and Ashton, for active 2003; anuran Silveira, species. 2005; In the case of P. fenestratus, there was no TheGriscom complexity et al., 2007) of the and structure being able of vegetation to influence of =0.04). this type of forest accounts for some species of Pristimantis to be very low. Fsignificant differencep of the hanger height P. altamazonicus distributed in the sampling units (ANOVA, 9,199 distribution=0.83, (R>0.59). p The structure of the cmIn diameter López-Rojas at breast et al. height (2015), with a closed canopy vegetation did2 not influence its vertical withwas detectable abundant inlitter, areas while with P. trees fenestratus greater thanprefers 30 p =0.27, = 0.13). Only leaf2 litter sites with bamboo and trees with a diameter at depth explained 51% of distribution (R =0,51, =0.02). Vol. 27 (1) 2018. 47-54 DOI: https://doi.org/10.24841/fa.v27i1.457 51 Vertical segregation in Pristimantis species from a bamboo forest in southeast of Amazonia, Brazil

P. skydmainus, one factors, that were not taken in this study, for a of the species most sensitive to bamboo, was better understanding of the vertical variation of brest height greater than 10. some cases individuals were observed at heights mostly restricted to heights of less than 45 cm, in microsimpatric species (syntopics). assemblage microhabitat of Pristimantis (Arroyo ACKNOWLEDGEMENTS higher than 200 cm concordant on the use of There was overlap in the location of the verticalet al., 2008). zone among some species, although they providing references. This work was supported differed in terms of abundance, ontogenetically Paulo Melo, for reading the manuscript and and in habitat distribution. The differentiation between adults and juveniles may be due to the by post-graduate fellowships from Coordenação fact that adults position themselves at strategic de Aperfeiçoamento de Pessoal em Nivel sites during the night in search of food and sexual AuthorizationSuperior (CAPES) and andBiodiversity Programa deInformation Pesquisa em Biodiversidade (PPbio) núcleo Acre to JJLR. position themselves for optimal vocalization sites,partners while (Duellman females, that and are Pyles, already 1983), gravid, males can assistanceSystem (SISBIO) in editing for this collecting manuscript. permits (license remain on the ground due to the increase in their number 37974-1). Thanks also to Paul Freed for

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